On the apterous line of the termite Velocitermes heteropterus (Isoptera: Termitidae): developmental pathways and cellulose digestion.

Termites are social insects with an extraordinary ability to digest cellulose. Termite societies are structured into castes, and patterns of postembryonic development vary between different termite species. The apterous line may exhibit polymorphism ("physical castes"), in which workers are dimorphic and soldiers can be either dimorphic or trimorphic. We examined the occurrence of polymorphism in the apterous line of Velocitermes heteropterus and determined the developmental pathways for this termite species. We also investigated the expression of the cellulase genes encoding ß-glucosidase and endo-ß-1,4-glucanase among the castes to determine whether there is a difference in digestion and, consequently, a possible division of labor with respect to this activity among the worker castes. The apterous line of V. heteropterus presents individuals of both sexes with two larval instars. The female larvae become major workers, and the male larvae become minor workers and soldiers. The expression of ß-glucosidase was similar within the castes, but the expression of endo-ß-1,4-glucanase was higher in workers than in soldiers. No significant differences were found between minor and major workers. These results suggest that there is no division of labor between the minors and majors with regard to cellulose digestion, with both workers contributing similarly to this process.

Labral gland in soldiers of the neotropical termite Cornitermes cumulans (Isoptera: Termitidae: Syntermitinae).

Cornitermes cumulans is a termite species of the subfamily Syntermitinae with a nasute mandibulate soldier caste, members of which are morphologically equipped with structures and organs specialized for the defence of the colony. We investigated the labrum of soldiers of C. cumulans and described the labral gland, an exocrine structure present in this appendage. The labrum of C. cumulans soldiers presented two distinct regions, the hyaline tip and the proximal region connected to the head. The hyaline tip exhibited a thick cuticle composed of a loose endocuticle involving an epithelium of class 1 cells, which synthesize the glandular product into a subcuticular space that function as a reservoir prior to release the final secretion. The proximal region of the labrum had an epithelium composed of class 1 and class 3 cells, which released the secretion onto both ventral and dorsal surfaces. The ultrastructure showed abundant smooth endoplasmic reticulum and glycogen in the class 1 cells, whereas the class 3 cells had many electron-lucent vesicles that varied in size. We associated the labral gland with the production of toxic substances that may act inside enemy wounds made by the soldiers' mandibles. Other possible functions for the labral secretion are also discussed.

Tergal glands in termite soldiers of the subfamily Syntermitinae (Isoptera: Termitidae).

The subfamily Syntermitinae comprises 14 genera of termites that are exclusively neotropical. The present study reports morphological data about mandibulate nasute soldiers from termite species belonging to three different genera within this subfamily. We describe tergal glands that were present under all tergites of soldiers of the following species: Cornitermes cumulans, Procornitermes araujoi, Syntermes nanus, and Syntermes wheeleri. The tergal glands were composed of class 2 and class 3 cells. Class 2 cells never reached the cuticle and were located below a flat layer of epidermal cells. Class 3 cells, composed of secretory cells and canal cells, were sporadic, whereas class 2 secretory cells were abundant. Secretory cells of class 3 were narrow and their cytoplasms were filled with several clear, oval-shaped vesicles with limiting membranes. The ultrastructure of class 2 cells showed well-developed smooth endoplasmic reticulum, Golgi apparatus, elongated mitochondria, several electron-lucent vesicles, and electron-dense granules that contain paracrystalline structures in S. nanus. Scanning electron micrographs displayed pores, campaniform sensilla and hairs in the outer cuticle of the soldier tergites. We hypothesize that soldier tergal glands may be involved in the production of defensive compounds, which occur in similar glands of certain cockroaches, or of primer pheromones, that might act in the regulation of soldier differentiation in the termite colony. To date, tergal glands have only been described in termite imagoes, and their occurrence in these soldiers of basal Syntermitinae implies a specific role in this caste that is still speculative and needs to be clarified.

Pheromones and exocrine glands in Isoptera.

Termites are eusocial insects that have a peculiar and intriguing system of communication using pheromones. The termite pheromones are composed of a blend of chemical substances and they coordinate different social interactions or activities, including foraging, building, mating, defense, and nestmate recognition. Some of these sociochemicals are volatile, spreading in the air, and others are contact pheromones, which are transmitted by trophallaxis and grooming. Among the termite semiochemicals, the most known are alarm, trail, sex pheromones, and hydrocarbons responsible for the recognition of nestmates. The sources of the pheromones are exocrine glands located all over the termite body. The principal exocrine structures considered pheromone-producing glands in Isoptera are the frontal, mandibular, salivary or labial, sternal, and tergal glands. The frontal gland is the source of alarm pheromone and defensive chemicals, but the mandibular secretions have been little studied and their function is not well established in Isoptera. The secretion of salivary glands involves numerous chemical compounds, some of them without pheromonal function. The worker saliva contains a phagostimulating pheromone and probably a building pheromone, while the salivary reservoir of some soldiers contains defensive chemicals. The sternal gland is the only source of trail-following pheromone, whereas sex pheromones are secreted by two glandular sources, the sternal and tergal glands. To date, the termite semiochemicals have indicated that few molecules are involved in their chemical communication, that is, the same compound may be secreted by different glands, different castes and species, and for different functions, depending on the concentration. In addition to the pheromonal parsimony, recent studies also indicate the occurrence of a synergic effect among the compounds involved in the chemical communication of Isoptera.